vuzbiochemiИзвестия вузов. Прикладная химия и биотехнологияProceedings of Universities. Applied Chemistry and Biotechnology2227-29252500-1558ИРНИТУ10.21285/2227-2925-2019-9-4-679-693vuzbiochemi-262Research ArticleФИЗИКО-ХИМИЧЕСКАЯ БИОЛОГИЯPHYSICOCHEMICAL BIOLOGYПроблемы и перспективы использования микроорганизмов для утилизации отходов лигноцеллюлозыProblems and prospects for the application of microorganisms in the disposal of lignocellulose wasteБолотниковаО. И.BolotnikovaO. I.

Болотникова Ольга Ивановна, к.б.н., доцент кафедры биомедицинской химии, иммунологии и лабораторной диагностики медицинского института

185910, г. Петрозаводск, пр-т Ленина, 33, Республика Карелия

Оlga I. Bolotnikova, Cand. Sci. (Biology), Associate Professor, Department of Biomedical Chemistry, Immunology and Laboratory Diagnostics, Institute of Medicine

33, Lenin Ave., Petrozavodsk 185910, Republic of Karelia

olga-bolotnikova@rambler.ruМихайловаН. П.MikhailovaN. P.

Михайлова Наталья Павловна, д.б.н., старший научный сотрудник кафедры молекулярной биотехнологии факультета химической и биотехнологии

190013, г. Санкт-Петербург, Московский пр-т, 26

Natalia P. Mikhailova, Dr. Sci. (Biology), Senior Researcher, Sub-Department of Molecular Biotechnology, Department Chemical and Biotechnology 

26, Moskovskii Ave., St. Petersburg 190013

m_natalia2@rambler.ruБазарноваЮ. Г.BazarnovaJu. G.

Базарнова Юлия Генриховна, д.т.н., профессор Высшей школы биотехнологии и пищевых технологий

195251, г. Санкт-Петербург, ул. Политехническая, 29

Julia G. Bazarnova, Dr. Sci. (Engineering), Professor, Graduate School of Biotechnology and Food Science

29, Politehnicheskaya St., St. Petersburg 195251

j.bazarnowa2012@yandex.ruАроноваЕ. Б.AronovaE. B.

Аронова Екатерина Борисовна, к.т.н., доцент Высшей школы биотехнологии и пищевых технологий

195251, г. Санкт-Петербург, ул. Политехническая, 29

Ekaterina B. Aronova, Cand. Sci. (Engineering), Associate Professor, Graduate School of Biotechnology and Food Science 

29, Politehnicheskaya St., St. Petersburg 195251,

aronovae@inbox.ruБолотниковаТ. А.BolotnikovaT. A.

Болотникова Татьяна Алексеевна, магистрант Высшей школы биотехнологии и пищевых технологий

195251, г. Санкт-Петербург, ул. Политехническая, 29

Тatyana А. Bolotnikova, Master Student, Graduate School of Biotechnology and Food Science

29, Politehnicheskaya St., St. Petersburg 195251

bolotnikova@ro.ruАкининаЮ. Н.AkininaJu. N.

Акинина Юлия Николаевна, магистрант кафедры физической химии факультета химии веществ и материалов

190013, г. Санкт-Петербург, Московский пр-т, 26

Julia N. Akinina, Master Student, Sub-Department of Physical Chemistry, Department of Substances and Materials Chemistry

26, Moskovskiy Ave., St. Petersburg 190013

akinina.ju@gmail.comПетрозаводский государственный университетРоссияPetrozavodsk State UniversityRussian FederationСанкт-Петербургский государственный технологический институт (технический университет)»РоссияSt. Petersburg State Institute of TechnologyRussian FederationСанкт-Петербургский политехнический университет Петра ВеликогоРоссияPeter the Great Saint-Petersburg Polytechnic UniversityRussian Federation20190501202094679693Copyright © Болотникова О.И., Михайлова Н.П., Базарнова Ю.Г., Аронова Е.Б., Болотникова Т.А., Акинина Ю.Н., 20202020Болотникова О.И., Михайлова Н.П., Базарнова Ю.Г., Аронова Е.Б., Болотникова Т.А., Акинина Ю.Н.Bolotnikova O.I., Mikhailova N.P., Bazarnova J.G., Aronova E.B., Bolotnikova T.A., Akinina J.N.Данная работа распространяется под лицензией Creative Commons Attribution 4.0.This work is licensed under a Creative Commons Attribution 4.0 License.https://vuzbiochemi.elpub.ru/jour/article/view/262

Конверсия бытовых и производственных отходов, содержащих лигноцеллюлозу, в разнообразные целевые продукты (источники биоэнергии, органические кислоты, сахарозаменители и т.д.) является одним из приоритетных направлений государственной экологической политики Российской Федерации. Однако рентабельность переработки субстратов, полученных в ходе гидролиза таких вторичных источников сырья, определяется возможностью микробиологической утилизации не только гексоз (D-глюкоза, D-манноза, D-галактоза), но и пентоз (D-ксилоза, L-арабиноза). Цель настоящего обзора – обсуждение перспектив использования микроорганизмов для утилизации пентоз лигноцеллюлозы, а также проблем, возникающих на пути технологической реализации этого процесса. В обзоре приведены современные данные о спектре про- и эукариотических микроорганизмов, обеспечивающих деструкцию лигноцеллюлозы и утилизацию ее структурных компонентов в природных экосистемах. Представлена краткая характеристика механизма действия ферментов лигниназного, целлюлазного и гемицеллюлазного комплексов. Выделены основные проблемы, сдерживающие применение энзиматического гидролиза многокомпонентных бытовых и промышленных отходов лигноцеллюлозы. Рассмотрены факторы, определяющие селективность катаболизма пентоз у мицелиальных грибов, бактерий и дрожжей. Определен спектр целевых продуктов биоконверсии пентоз лигноцеллюлозы, имеющих народно-хозяйственную значимость. Обсуждаются способы комплексной микробиологической утилизации разнообразных бытовых и сельскохозяйственных отходов, а также возможность вовлечения в данный процесс побочных продуктов промышленной деструкции древесины (кислотных гидролизатов и сульфитных щелоков).

Авторы заявляют об отсутствии конфликта интересов.

The conversion of household and industrial wastes containing lignocellulose into a variety of target products (bioenergy sources, organic acids, sweeteners, etc.) involves one of the priority directions for the state environmental policy of the Russian Federation. However, the profitability of processing the substrates obtained by hydrolysis of such secondary sources of raw materials is determined by the possibility of microbiological utilisation for not only hexoses (D-glucose, D-mannose, D-galactose), but also pentose (D-xylose, L-arabinose). The aim of this review consists in a discussion of the prospects for using microorganisms in the disposal of lignocellulose pentoses, along with problems arising in the course of the technological implementation of this process. The review provides contemporary data on the spectrum of pro- and eukaryotic microorganisms ensuring the destruction of lignocellulose and the utilisation of its structural components in natural ecosystems. A brief description of action mechanism inherited to the enzymes of ligninase, cellulase and hemicellulase complexes is presented. The main problems hindering the enzymatic hydrolysis application to multicomponent household and industrial lignocellulose wastes are identified. The factors determining the selectivity of pentosis catabolism in mycelial fungi, bacteria and yeast are examined. The spectrum of target products in bioconversion of lignocellulose pentoses, is determined with regard of their economic importance. The methods of complex microbiological utilisation of various household and agricultural wastes, as well as the possibility of involving by-products from industrial destruction of wood (acid hydrolysates and sulphite liquors) in this process, are discussed.

The authors declare no conflict of interests regarding the publication of this article.

отходы лигноцеллюлозыD-ксилозаL-арабинозабиоконверсиягрибыбактерииlignocellulose wasteD-xyloseL-arabinosebioconversionfungibacteriaНаучный обзор подготовлен при поддержке гранта РФФИ № 18-44-100001.The scientific review was supported by grant 18-44-100001 from Russian Foundation for Basic Research.ReferencesКальнер В.Д. Экологически ориентированная среда обитания - интегральный критерий качества жизни // Экология и промышленность России. 2019. Т. 23. N 10. С. 50–55.Kal'ner VD. An ecologically oriented human environment is an integral criterion for the quality of life. Ekologiya i promyshlennost' Rossii = Ecology and Industry of Russia. 2019;23(10):50–55. (In Russian)Болотникова О.И., МихайловаН.П., Гинак А.И. Кислотный и энзиматический гидролиз непищевых источников растительной биомассы: перспективы промышленной реализации (обзор) // Известия Санкт-Петербургского государственного технологического института (технического университета). 2017. N 39 (65). С. 89–95.Bolotnikova OI, Mikhailova NP, Ginak AI. Acid and enzymatic hydrolysis of non- food-based biomass sources: prospects for industrial implementation. Izvestiya Sankt-Peterburgskogo gosudarstvennogo tekhnologicheskogo instituta (tekhnicheskogo universiteta). = Bulletin of the Saint Petersburg State Institute of Technology (Technical University). 2017;39:89–95. (In Russian)Cheng H., Wang L. Lignocelluloses feedstock biorefinery as petrorefinery substitutes. Open access peer-reviewed chapter. 2013. Available from: https://www.intechopen.com/books/biomassnow-sustainable-growth-and-use/lignocellulosesfeedstock-biorefinery-as-petrorefinery-substitutes [Accessed 26th Januaru 2019].Cheng H, Wang L. Lignocelluloses feedstock biorefinery as petrorefinery substitutes. Open access peer-reviewed chapter. 2013. Available from: https://www.intechopen.com/books/biomass-nowsustainable-growth-and-use/lignocelluloses-feed stock-biorefinery-as-petrorefinery-substitutes [Accessed 26th Januaru 2019].Chen H. Biotechnology of lignocellulose, theory and practice. Chemical Industry Press, Beijing and Springer, 2014. 510 p.Chen H. Biotechnology of lignocellulose, theory and practice. Chemical Industry Press, Beijing and Springer; 2014. 510 p.Гунич С.В., Янчуковская Е.В., Днепровская Н.И. Анализ современных методов переработки твердых бытовых отходов // Известия вузов. Прикладная химия и биотехнология. 2015. N 2 (13). С. 110–115.Gunich SV, Yanchukovskaya EV, Dneprovskaya NI. Analysis of modern methods of hard domestic wastes processing. Izvestiya Vuzov. Prikladnaya Khimiya i Biotekhnologiya = Proceedings of Universities. Applied Chemistry and Biotechnology. 2015;2:110–115. (In Russian)Щукина Т.В. Биогаз – перспективы и возможности производства // Известия вузов. Прикладная химия и биотехнология. 2012. N 1 (2). С. 113–118.Shchukina TV. Biogas – prospects and manufacture possibilities. Izvestiya Vuzov. Prikladnaya Khimiya i Biotekhnologiya = Proceedings of Universities. Applied Chemistry and Biotechnology. 2012;1: 113–118. (In Russian)Чхенкели В.А., Глушенкова Т.В., Горяева Н.А., Чхенкели Л.Г., Калинович А.Е. Оптимизация получения биологически активной субстанции на основе гриба-ксилотрофа Trametes pubescens shumach.: Fr.)Pilat // Известия вузов. Прикладная химия и биотехнология. 2011. N 1 (1). С. 84–89.ChkhenkeliVA, GlushenkovaTV, Goryaeva NA, Chkhenkeli LG, Kalinovich AE. Optimization of production of biologically active substance on the basis of the fungus-xylotroph Trametes pubescens shumach.: Fr.)Pilat. Izvestiya Vuzov. Prikladnaya Khimiya i Biotekhnologiya = Proceedings of Universities. Applied Chemistry and Biotechnology. 2011;1:84–89. (In Russian)Maitan-Alfenas G.P., Visser E.M., Guimarães V.M. Enzymatic hydrolysis of lignocellulosic biomass: converting food waste in valuable products // Current Opinion in Food Science. 2015. Vol. 1. P. 44–49. https://doi.org/10.1016/j.cofs.2014.10.001Maitan-Alfenas GP, Visser EM, Guimarães VM. Enzymatic hydrolysis of lignocellulosic bio mass: converting food waste in valuable products. Current Opinion in Food Science. 2015;1:44–49. https://doi.org/10.1016/j.cofs.2014.10.001Ferreira J.A., Mahboubi A., Lennartsson P.R., Taherzadeh M.J. Waste biorefineries using filamentous ascomycetes fungi: present status and future prospects // Bioresource Technology. 2016. Vol. 215. P. 334–345. https://doi.org/10.1016/j.biortech.2016.03.018Ferreira JA, Mahboubi A, Lennartsson PR, Taherzadeh MJ. Waste biorefineries using filamentous ascomycetes fungi: present status and future prospects. Bioresource Technology. 2016;215:334– 345. https://doi.org/10.1016/j.biortech.2016.03.018Kulikova N.A., Kleina O.I., Stepanova E.V., Koroleva O.V. Use of Basidiomycetes in Industrial Waste Processing and Utilization Technologies: Fundamental and Applied Aspects (Review) // Applied Biochemistry and Microbiology. 2011. Т. 47. N 6. С. 619–634.Kulikova NA, Kleina OI, Stepanova EV, Koroleva OV. Use of Basidiomycetes in Industrial Waste Processing and Utilization Technologies: Fundamental and Applied Aspects (Review). Applied Biochemistry and Microbiology. 2011;47(6): 619–634.Jönsson L.J., Alriksson B., Nilvebrant N.-O. Bioconversion of lignocellulose: inhibitors and detoxification // Biotechnology for Biofuels. 2013. Vol. 6. Issue 1. P. 16–26. https://doi.org/10.1186/1754-6834-6-16Jönsson LJ, Alriksson B, Nilvebrant N-O. Bioconversion of lignocellulose: inhibitors and detoxification. Biotechnology for Biofuels. 2013;6(1): 16–26. https://doi.org/10.1186/1754-6834-6-16Górska E.B., Jankiewicz U., Dobrzyński J., Gałązka А., Sitarek M., Gozdowski D., et al. Production of ligninolytic enzymes by cultures of white rot fungi // Polish journal of microbiology / Polskie Towarzystwo Mikrobiologów = The Polish Society of Microbiologists. 2014. Vol. 63. Issue 4. P. 461–465. https://doi.org/10.33073/pjm-2014-062Górska EB, Jankiewicz U, Dobrzyński J, Gałązka А, Sitarek M, Gozdowski D, et al. Production of ligninolytic enzymes by cultures of white rot fungi. Polish journal of microbiology. 2014;63(4): 461–465. https://doi.org/10.33073/pjm-2014-062Plácido J., Capareda S. Ligninolytic enzymes: a biotechnological alternative for bioethanol production // Bioresources and Bioprocessing. 2015. Vol. 2. 12 p. https://doi.org/10.1186/s40643- 015-0049-5Plácido J, Capareda S. Ligninolytic enzymes: a biotechnological alternative for bioethanol production. Bioresources and Bioprocessing. 2015;2: 12 p. https://doi.org/10.1186/s40643-015-0049-5Sokan-Adeaga A.A., Ana Godson R.E.E., Sokan-Adeaga M.A., Sokan-Adeaga E.D. Lignocelluloses: An economical and ecological resource for bio-ethanol production. A Review // International Journal of Natural Resource Ecology and Management. 2016. Vol. 1. Issue 3. p. 128–144. https://doi.org/10.11648/j.ijnrem.20160103.18Sokan-Adeaga AA, Ana Godson REE, SokanAdeaga MA, Sokan-Adeaga ED. Lignocelluloses: An economical and ecological resource for bioethanol production. A Review. International Journal of Natural Resource Ecology and Management. 2016;1(3):128–144. https://doi.org/10.11648/j.ijnrem. 20160103.18Приставка А.А., Попова И.В. Влияние фторида натрия на ферментативную активность грибных целлюлаз // Известия вузов. Прикладная химия и биотехнология. 2015. N 1 (12). С. 36–46.Pristavka AA, Popova IV. Influence of sodium fluoride on enzymatic activity of fungal cellulases. Izvestiya Vuzov. Prikladnaya Khimiya i Biotekhnologiya = Proceedings of Universities. Applied Chemistry and Biotechnology. 2015;1:36–46. (In Russian)Gavrilov S.N., Stracke C., Jensen K., Menzel P., Kallnik V., Slesarev A., et al. Isolation and characterization of the first xylanolytic hyperthermophilic euryarchaeon Thermococcus sp. strain 2319×1 and its unusual multidomain glycosidase // Frontiers in Microbiology. 2016. Vol. 7. Issue 75. P. 552–569. https://doi.org/10.3389/fmicb.2016.00552Gavrilov SN, Stracke C, Jensen K, Menzel P, Kallnik V, Slesarev A, et al. Isolation and characterization of the first xylanolytic hyperthermophilic euryarchaeon Thermococcus sp. strain 2319×1 and its unusual multidomain glycosidase. Frontiers in Microbiology. 2016;7(75):552–569. https://doi.org/10.3389/fmicb.2016.00552Peng X., Qiao W., Mi S., Jia X., Su H., Han Y. Characterization of hemicellulase and cellulase from the extremely thermophilic bacterium Caldicellulosiruptor owensensis and their potential application for bioconversion of lignocellulosic biomass without pretreatment // Biotechnology for Biofuels. 2015. Vol. 8. Issue 1. P. 131–145. https://doi.org/10.1186/s 13068-015-0313-0Peng X, Qiao W, Mi S, Jia X, Su H, Han Y. Characterization of hemicellulase and cellulase from the extremely thermophilic bacterium Caldicellulosiruptor owensensis and their potential application for bioconversion of lignocellulosic biomass without pretreatment. Biotechnology for Biofuels. 2015;8(1): 131–145. https://doi.org/10.1186/s13068-015-0313-0KurtzmanC.P., Fell J.W., Boekhout T. (eds.). The Yeasts A Taxonomic Study, 5th Edition. Elsevier, Amsterdam, Netherlands. 2011. 2354 p. https://doi.org/10.1016/B978-0-444-52149-1.00007-0Kurtzman CP, Fell JW, Boekhout T. (eds.). The Yeasts A Taxonomic Study. 5th ed. Elsevier, Amsterdam, Netherlands; 2011. 2354 p. https://doi.org/10.1016/B978-0-444-52149-1.00007-0Mahboubi A., Ferreira J.A., Taherzadeh M.J., Lennartsson P.R. Production of fungal biomass for feed, fatty acids, and glycerol by Aspergillus oryzae from fat-rich dairy substrates // Fermentation. 2017. Vol. 3. Issue 4. P. 48–58. https://doi.org/10.3390/fermentation3040048Mahboubi A, Ferreira JA, Taherzadeh MJ, Lennartsson PR. Production of fungal biomass for feed, fatty acids, and glycerol by Aspergillus oryzae from fat-rich dairy substrates. Fermentation. 2017;3(4): 48–58. https://doi.org/10.3390/fermentation3040048Bolotnikova O.I., Trushnikova E.P., Mikhailova N.P., Ginak A.I. Production of xylitol and ethanol and activity of the key enzymes of D-xylose consumption in Pachysolen tannophilus mutant strains // Microbiology. 2015. Vol. 84. N 4. P. 479–484. https://doi.org/10.1134/S0026261715040049Bolotnikova OI, Trushnikova EP, Mikhailova NP, Ginak AI. Production of xylitol and ethanol and activity of the key enzymes of D-xylose consumption in Pachysolen tannophilus mutant strains. Microbiology. 2015;84(4):479–484. https://doi.org/10. 1134/S0026261715040049De Groot M.J., Prathumpai W., Visser J., Ruijter G.J. Metabolic control analysis of Aspergillus niger L-arabinose catabolism // Biotechnology Progress. 2005. Vol. 21. Issue 6. P. 1610–1616. https://doi.org/10.1021/bp050189oDe Groot MJ, Prathumpai W, Visser J, Ruijter GJ. Metabolic control analysis of Aspergillus niger L-arabinose catabolism. Biotechnology Progress. 2005; 21(6):1610–1616. https://doi.org/10.1021/bp050189oKoirala S., Wang X., Rao C.V. Reciprocal regulation of L-arabinose and D-xylose metabolism in Escherichia coli // Journal of Bacteriology. 2016. Vol. 198. Issue 3. P. 386–393. https://doi.org/10. 1128/JB.00709-15Koirala S, Wang X, Rao CV. Reciprocal regulation of L-arabinose and D-xylose metabolism in Escherichia coli. Journal of Bacteriology. 2016; 198(3):386–393. https://doi.org/10.1128/JB.00709-15Chen X., Jiang Z.-H., Chen S., Qin W. Microbial and bioconversion production of D-xylitol and its detection and application // International Journal of Biological Sciences. 2010. Vol. 6. Issue 7. P. 834–844. https://doi.org/10.7150/ijbs.6.834Chen X, Jiang Z-H, Chen S, Qin W. Microbial and bioconversion production of D-xylitol and its detection and application. International Journal of Biological Sciences. 2010;6(7):834–844. https://doi.org/10.7150/ijbs.6.834Saha B.C., Kennedy G.J., Qureshi N., Bowman M.J. Production of itaconic acid from pentose sugars by Aspergillus terreus // Biotechnology Progress. 2017. Vol. 33. Issue 4. P. 1059–1067. https://doi.org/10.1002/btpr.2485Saha BC, Kennedy GJ, Qureshi N, Bowman MJ. Production of itaconic acid from pentose sugars by Aspergillus terreus. Biotechnology Progress. 2017;33(4):1059–1067. https://doi.org/10.10 02/btpr.2485Watanabe S., Kodaki T., Makino K. L-Arabinose 1-dehydrogenase: A novel enzyme involving in bacterial L-arabinose metabolism // Nucleic Acids Symposium Series. 2005. Vol.49. Issue 1. P. 309–310. https://doi.org/10.1093/nass/49.1.309Watanabe S, Kodaki T, Makino K. L-Arabinose 1-dehydrogenase: A novel enzyme involving in bacterial L-arabinose metabolism. Nucleic Acids Symposium Series. 2005;49(1):309–310. https://doi.org/10.1093/nass/49.1.309Bajerski F., Ganzert L., Mangelsdorf K., Lipski A., Busse H.-J., Padur L., et al. Herbaspirillum psychrotolerans sp. nov., a member of the family Oxalobacteraceae from a glacier forefield // International Journal of Systematic and Evolutionary Microbiology. 2013. Vol. 63. Issue 9. P. 3197–3203. https://doi.org/10.1099/ijs.0.046920-0Bajerski F, Ganzert L, Mangelsdorf K, Lipski A, Busse H-J, Padur L, et al. Herbaspirillum psychrotolerans sp. nov., a member of the family Oxalobacteraceae from a glacier forefield. International Journal of Systematic and Evolutionary Microbiology. 2013;63(9):3197–3203. https://doi.org/10.1099/ijs.0.046920-0Zhao B., Chen S. Alkalitalea saponilacus gen.nov., sp. nov., an obligately anaerobic, alkaliphilic, xylanolytic bacterium from a meromictic soda lake // International Journal of Systematic and Evolutionary Microbiology. 2012. Vol. 62. Issue 11. P. 2618–2623. https://doi.org/10.1099/ijs.0.038315-0Zhao B, Chen S. Alkalitalea saponilacus gen. nov., sp. nov., an obligately anaerobic, alkaliphilic, xylanolytic bacterium from a meromictic soda lake. International Journal of Systematic and Evolutionary Microbiology. 2012;62(11):2618–2623. https://doi.org/10.1099/ijs.0.038315-0Zhilina T.N., Kevbrin V.V., Tourova T., Lysenko A.M., Kostrikina N.A., Zavarzin G.A. Clostridium alkalicellum sp. nov., an obligately alkaliphilic cellulolytic bacterium from a soda lake in the baikal region // Microbiology. 2005. Vol. 74. Issue 5. P. 557–566. https://doi.org/10.1007/s11021-005-0103-yZhilina TN, Kevbrin VV, Tourova T, Lysenko AM, Kostrikina NA, Zavarzin GA. Clostridium alkalicellum sp. nov., an obligately alkaliphilic cellulolytic bacterium from a soda lake in the baikal region. Microbiology. 2005;74(5):557–566. https://doi.org/10.1007/s11021-005-0103-yPugin B., Blamey J.M., Baxter B.K., Wiegel J. Amphibacillus cookii sp. nov., a facultatively aerobic, spore-forming, moderately halophilic, alkalithermotolerant bacterium // International Journal of Systematic and Evolutionary Microbiology. 2012. Vol. 62. Issue 9. P. 2090–2096. https://doi.org/10. 1099/ijs.0.034629-0Pugin B, Blamey JM, Baxter BK, Wiegel J. Amphibacillus cookii sp. nov., a facultatively aerobic, spore-forming, moderately halophilic, alkalithermotolerant bacterium. International Journal of Systematic and Evolutionary Microbiology. 2012;62(9): 2090–2096. https://doi.org/10.1099/ijs.0.034629-0Choudhury D., Gayen K., Saini S. Dynamic control of arabinose and xylose utilization in E. coli // The Canadian Journal of Chemical Engineering. 2018. Vol. 96. Issue 9. P. 1881–1887. https://doi.org/10.1002/cjce.23197Luo H, Wu Y, Kole C. Compendium of Bioenergy Plants: Switchgrass. CRC Press; 2014. 464 p.Luo H., Wu Y., Kole C. Compendium of Bioenergy Plants: Switchgrass. CRC Press. 2014. 464 p.Zimmermann FK, Entian K-D. Yeast Sugar Metabolism. Biochemistry, Genetics, Biotechnology and Application. CRC Press; 1997. 567 p.ZimmermannF.K., Entian K.-D. Yeast Sugar Metabolism. Biochemistry, Genetics, Biotechnology and Application. CRC Press, 1997. 567 p.Gupta R, Mehta G, Kuhad RC. Fermentation of pentose and hexose sugars from corncob, a low cost feedstock into ethanol. Biomass and Bioenergy. 2012;47:334–341. https://doi.org/10.1016/j.biombioe. 2012.09.027Gupta R., Mehta G., Kuhad R.C. Fermentation of pentose and hexose sugars from corncob, a low cost feedstock into ethanol // Biomass and Bioenergy. 2012. Vol. 47. P. 334–341. https://doi.org/10. 1016/j.biombioe.2012.09.027Skiba EA, Mironova GF. Advantages of combining biocatalytic stages in bioethanol synthesis from cellulosic biomasses. Izvestiya Vuzov. Prikladnaya Khimiya i Biotekhnologiya = Proceedings of Higher School. Applied Chemistry and Biotechnology. 2016;6(4):53–60. (In Russian) https://doi.org/10. 21285/2227-2925-2016-6- 4-53-60Скиба Е.А., Миронова Г.Ф. Преимущества совмещения биокаталических стадий в синтезе биоэтанола из целлюлозосодержащего сырья // Известия вузов. Прикладная химия и биотехнология. 2016. Т. 6. N 4. С. 53–60. https://doi.org/10.21285/2227-2925-2016-6- 4-53-60Kozlov IA, Garipov RM. Catalysis and its impact in the processes of deep processing of plant biomass. Izvestiya Vuzov. Prikladnaya Khimiya i Biotekhnologiya = Proceedings of Higher School. Applied Chemistry and Biotechnology. 2017;7(1):188–191. (In Russian) https://doi.org/10.21285/2227-2925-2017- 7-1-188-191Козлов И.А., Гарипов Р.М. Катализ и его роль в процессах глубокой переработки растительной биомассы // Известия вузов. Прикладная химия и биотехнология. 2017. Т. 7. N 1. С. 188–191. https://doi.org/10.21285/2227-2925- 2017-7-1-188-191Makarova E.I., Budaeva V.V. Estimation of the efficiency of the oat bran enzymatic hydrolysis with feeding at high initial substrate concentrations. Izvestiya Vuzov. Prikladnaya Khimiya i Biotekhnologiya = Proceedings of Higher School. Applied Chemistry and Biotechnology. 2017:7(4):51–57. (In Russian) https://doi.org/10.21285/2227-2925-2017-7- 4-51-57Макарова Е.И., Будаева В.В. Оценка эффективности ферментативного гидролиза плодовых оболочек овса с подпиткой при высоких начальных концентрациях субстрата // Известия вузов. Прикладная химия и биотехнология. 2017. Т.7. N4. С.51–57. https://doi.org/10.21285/2227- 2925-2017-7-4-51-57Molokova KV, Privalova EA, Gil TA. Cultivation of clostridium acetobutylicum vkm 1787 – producer of butanol, acetone and ethanol. Izvestiya Vuzov. Prikladnaya Khimiya i Biotekhnologiya = Proceedings of Higher School. Applied Chemistry and Biotechnology. 2013:1(4):87–91. (In Russian)Молокова К.В., Привалова Е.А., Гиль Т.А. Культивирование Clostridium acetobutylicum ВКМ 1787 – продуцента бутанола, ацетон а и этанола. // Известия вузов. Прикладная химия и биотехнология. 2013. N 1 (4). С. 87–91.Raganati F, Olivieri G, Russo ME, Marzocchella A. Butanol production by Clostridium acetobutylicum in a continuous packed bed reactor fed with cheese whey. Chemical Engineering Transactions. 2013;32:937–642. https://doi.org/10.3303/CET1332157Raganati F., Olivieri G., Russo M.E., Marzocchella A. Butanol production by Clostridium acetobutylicum in a continuous packed bed reactor fed with cheese whey // Chemical Engineering Transactions. 2013. Vol. 32. P. 937–642. https://doi.org/10.3303/CET1332157Sousa JAB, Sorokin DY, Bijmans MFM, Plugge CM, Stams AJM. Ecology and application of haloalkaliphilic anaerobic microbial communities. Applied Microbiology and Biotechnology. 2015; 99(22): 9331–9336. https://doi.org/10.1007/s00253-015-6937-ySousa J.A.B., Sorokin D.Y., Bijmans M.F.M., Plugge C.M., Stams A.J.M. Ecology and application of haloalkaliphilic anaerobic microbial communities // Applied Microbiology and Biotechnology. 2015. Vol. 99. Issue 22. P. 9331–9336. https://doi.org/10.1007/s00253-015-6937-yJung M-Y, Mazumdar S, Shin SH, Yang KS, Lee J, Oh M-K. Improvement of 2,3-butanediol yield in Klebsiella pneumoniae by deletion of the pyruvate formate-lyase gene. Applied and Environmental Microbiology. 2014;80(19):6195–6203. https://doi.org/10.1128/AEM.02069-14JungM.-Y., MazumdarS., ShinS.H., Yang K.-S., Lee J., Oh M.-K. Improvement of 2,3-butanediol yield in Klebsiella pneumoniae by deletion of the pyruvate formate-lyase gene // Applied and Environmental Microbiology. 2014. Vol. 80. Issue 19. P. 6195–6203. https://doi.org/10.1128/AEM.02069-14Gupta A, Murarka A, Campbell P, Gonzalez R. Anaerobic fermentation of glycerol in paenibacillus macerans: Metabolic pathways and environmental determinants. Applied and Environmental Microbiology. 2009;75.(18):5871–5883. http://dx.doi.org/10.1128/AEM.01246-09Gupta A., Murarka A., Campbell P., Gonzalez R. Anaerobic fermentation of glycerol in paenibacillus macerans: Metabolic pathways and environmental determinants // Applied and Environmental Microbiology. 2009. Vol. 75. Issue 18. P. 5871– 5883. http://dx.doi.org/10.1128/AEM.01246-09Sravanthi T, Tushar L, Sasikala Ch, Ramana ChV. Alkalispirochaeta cellulosivorans gen. nov., sp. nov., a cellulose-hydrolysing, alkaliphilic, halotolerant bacterium isolated from the gut of a woodeating cockroach (Cryptocercus punctulatus), and reclassification of four species of Spirochaeta as new combinations within Alkalispirochaeta gen. nov. International Journal of Systematic and Evolutionary Microbiology. 2016;66(4):1612–1619. https://doi.org/10.1099/ijsem.0.000865SravanthiT.,Tushar L., Sasikala Ch., Ramana Ch.V. Alkalispirochaeta cellulosivorans gen. nov., sp. nov., a cellulose-hydrolysing, alkaliphilic, halotolerant bacterium isolated from the gut of a woodeating cockroach (Cryptocercus punctulatus), and reclassification of four species of Spirochaeta as new combinations within Alkalispirochaeta gen. nov. // Inter-national Journal of Systematic and Evolutionary Microbiology. 2016. Vol. 66. Issue 4. P. 1612–1619. https://doi.org/10.1099/ijsem.0.000865Balasubramanian N, Kim JS, Lee YY. Fermentation of xylose into acetic acid by Clostridium thermoaceticum. Applied Biochemistry and Biotechnology. 2001;91(1-9):367–376. https://doi.org/10.1385/abab:91-93:1-9:367Balasubramanian N., Kim J.S., Lee Y.Y. Fermentation of xylose into acetic acid by Clostridium thermoaceticum // Applied Biochemistry and Biotechnology. 2001. Vol. 91. Issue 1-9. P. 367– 376. https://doi.org/10.1385/abab:91-93:1-9:367Liu H, Wang W, Deng L, Wang F, Tan T. High production of fumaric acid from xylose by newly selected strain Rhizopus arrhizus RH 7-13-9#. Bioresource Technology. 2015;186:348–350. https://doi.org/10.1016/j.biortech.2015.03.109Liu H., Wang W., Deng L., Wang F., Tan T. High production of fumaric acid from xylose by newly selected strain Rhizopus arrhizus RH 7-13-9# // Bioresource Technology. 2015. Vol. 186. P. 348–350. https://doi.org/10.1016/j.biortech.2015.03.109Anasontzis GE, Christakopoulos P. Challenges in ethanol production with Fusarium oxysporum through consolidated bioprocessing. Bioengineered. 2014;5(6):393–395. https://doi.org/10.4161/bioe.36328Anasontzis G.E, Christakopoulos P. Challenges in ethanol production with Fusarium oxysporum through consolidated bioprocessing // Bioengineered. 2014. Vol. 5. Issue 6. P. 393–395. https://doi.org/10.4161/bioe.36328Assis LF, Kagohara E, Omori ÁT, Comasseto JV, Andrade LH, Porto ALM. Deracemization of (RS)-1-[(4-methylselanyl)phenyl]ethanol and (rs)-1- [(4-ethylselanyl)phenyl]ethanol by strains of Aspergillus terreus. Food Technology and Biotechnology. 2007;45(4):415–419.Assis L.F., Kagohara E., Omori Á.T., Comasseto J.V., Andrade L.H., Porto A.L.M. Deracemization of (RS)-1-[(4-methylselanyl)phenyl]ethanol and (rs)-1-[(4-ethylselanyl)phenyl]ethanol by strains of Aspergillus terreus // Food Technology and Biotechnology. 2007. Vol. 45. Issue 4. P. 415–419.Chandel AK, Kapoor RK, Singh A, Kuhad RC. Detoxification of sugarcane bagasse hydrolysate improves ethanol production by Candida shehatae NCIM 3501. Bioresource Technology. 2007;98(10): 1947–1950. https://doi.org/10.1016/j.biortech.2006.07.047Chandel A.K., Kapoor R.K., Singh A., Kuhad R.C. Detoxification of sugarcane bagasse hydrolysate improves ethanol production by Candida shehatae NCIM 3501 // Bioresource Technology. 2007. Vol. 98. Issue 10. P. 1947–1950. https://doi.org/10.1016/j.biortech.2006.07.047Agbogbo FK, Coward-Kelly G. Cellulosic ethanol production using the naturally occurring xylosefermenting yeast, Pichia stipites. Biotechnology Letters. 2008;30(9):1515–1524. https://doi.org/10.1007/s10529-008-9728-zAgbogbo F.K., Coward-Kelly G. Cellulosic ethanol production using the naturally occurring xylose-fermenting yeast, Pichia stipitis // Biotechnology Letters. 2008. Vol. 30. Issue 9. P. 1515–1524. https://doi.org/10.1007/s10529-008-9728-zPrakash G, Varma AJ, Prabhune A, Shouche Y, Rao M. Microbial production of xylitol from D-xylose and sugarcane bagasse hemicellulose using newly isolated thermotolerant yeast Debaryomyces hansenii. Bioresource Technology. 2011;102(3): 3304–3308. https://doi.org/10.1016/j.biortech.2010.10.074Prakash G., Varma A.J., Prabhune A., Shouche Y., Rao M. Microbial production of xylitol from D-xylose and sugarcane bagasse hemicellulose using newly isolated thermotolerant yeast Debaryomyces hansenii // Bioresource Technology. 2011. Vol. 102. Issue 3. P. 3304–3308. https://doi.org/10.1016/j.biortech.2010.10.074Zou YZ, Qi K, Chen X, Miao XL, Zhong JJ. Favorable effect of very low initial KLa value on xylitol production from xylose by a self-isolated strain of Pichia guilliermondii. Journal of Bioscience and Bioengineering. 2010;109(2):149–152. https://doi.org/10. 1016/j.jbiosc.2009.07.013ZouY.Z., QiK., ChenX., MiaoX.L., Zhong J.J. Favorable effect of very low initial KLa value on xylitol production from xylose by a self-isolated strain of Pichia guilliermondii // Journal of Bioscience and Bioengineering. 2010. Vol. 109. Issue 2. P. 149–152. https://doi.org/10.1016/j.jbiosc.2009.07.013Dashtban M, Schraft H, Qin W. Fungal bioconversion of lignocellulosic residues; Opportunities & Perspectives. International Journal of Biological Sciences. 2009:5(6):578–595. https://doi.org/10.7150/ijbs.5.578Dashtban M., Schraft H., Qin W. Fungal bioconversion of lignocellulosic residues; Opportunities & Perspectives // International Journal of Biological Sciences. 2009. Vol. 5. Issue 6. P. 578–595. https://doi.org/10.7150/ijbs.5.578Vazetdinova AA, Kharina MV, Loginova IV, Kleschevnikov LI. Enzymatic hydrolysis of cellulosic residuals of furfural production from vegetable raw materials. Bashkirskii khimicheskii zhurnal = Bashkir Chemical Journal. 2017;24(1):27–31. (In Russian)Вазетдинова А.А., Харина М.В., Логинова И.В., Клещевников Л.И. Ферментолиз целлюлозосодержащих остатков производства фурфурола из отходов растительного сырья // Башкирский химический журнал. 2017. Т.24. N 1. С. 27–31.Kharina MV, Grigor'eva ON. Design features of reactors for acid hydrolysis of lignocellulosecontaining raw materials. Vestnik Tekhnologicheskogo universiteta = Bulletin of the Technological University. 2017;20(13):143–150. (In Russian)Харина М.В., Григорьева О.Н. Особенности конструкции реакторов для кислотного гидролиза лигноцеллюлозосодержащего сырья // Вестник Технологического университета. 2017. Т. 20. N 13. С. 143–150.Fazliev II, Minzanova ST, Akhmadullina F Yu, Mukhachev SG. The effect of different acids on grain hydrolysis. Izvestiya vuzov. Prikladnaya khimiya i biotekhnologiya = Proceedings of Higher School. Applied Chemistry and Biotechnology. 2012;2:50–53. (In Russian)Фазлиев И.И., Минзанова С.Т., Ахмадуллина Ф.Ю., Мухачев С.Г. Влияние кислот различной природы на гидролиз пивной дробины // Известия вузов. Прикладная химия и биотехнология. 2012. N 2 (3). С. 50–53.Grigor'eva ON, Kharina MV. Acid hydrolysis of lignocellulose-containing raw materials in bioethanol production technology. Vestnik Tekhnologicheskogo universiteta = Bulletin of the Technological University. 2016;19(10):128–132. (In Russian)Григорьева О.Н., Харина М.В. Кислотный гидролиз лигноцеллюлозосодержащего сырья в технологии получения биоэтанола // Вестник Технологического университета. 2016. Т. 19. N 10. С. 128–132.Loginova IV, Kharina MV. Study of hightemperature autohydrolysis of lignocellulosic raw materials. Vestnik Tekhnologicheskogo universiteta = Bulletin of the Technological University. 2017;20(6): 143–145. (In Russian)Логинова И.В., Харина М.В. Исследование высокотемпературного автогидролиза лигноцеллюлозного сырья // Вестник Технологического университета. 2017. Т. 20. N 6. С. 143–145.Bolotnikova OI, Mikhailova NP, Ginak AI. Comparative analysis of the growth physiology of xylose-assimilating yeasts candida shehtae and pachysolen tannophilus. Mikologiya i fitopatologiya = Mycology and Phytopathology. 2013;47(5):329–332. (In Russian)Болотникова О.И., Михайлова Н.П., Гинак А.И. Сравнительный анализ физиологии роста ксилозоассимилирующих дрожжей Candida shehatae и Pachysolen tannophilus // Микология и фитопатология. 2013. Т. 47. N 5. С. 329–332.Болотникова О.И., Михайлова Н.П., Гинак А.И. Сравнительный анализ физиологии роста ксилозоассимилирующих дрожжей Candida shehatae и Pachysolen tannophilus // Микология и фитопатология. 2013. Т. 47. N 5. С. 329–332.

The authors declare that there are no conflicts of interest present.